­­­Two new species of Collembola (Hexapoda) from Saliente Cave (Almería, Spain)

Rafael JORDANA University of Navarra, Faculty of Sciences, Department of Environmental Biology, University Campus, 31080 Pamplona (Spain) [email protected] (corresponding author)

Pablo BARRANCO Department of Biology and Geology, Engineering High School, University of Almería. Cite-IIB, 04120 Almería (Spain) [email protected]

Ana AMEZCUA Enrique BAQUERO University of Navarra, Faculty of Sciences, Department of Environmental Biology, University Campus, 31080 Pamplona (Spain) [email protected]

Published on 31st March 2017

urn:lsid:zoobank.org:pub:C2ABEE8B-F4BB-46D7-9BE9-4E5E425C9CB7

Jordana R., Barranco P., Amezcua A. & Baquero E. 2017. — ­­­Two new species of Collembola (Hexapoda) from Saliente Cave (Almería, Spain). Zoosystema 39 (1): 103-115. https://doi.org/10.5252/z2017n1a12

ABSTRACT Two new species of cavernicolous Collembola belonging to genera Vargovitsh, 2009 and Pseudosinella Schäffer, 1897 from “Cueva del Saliente” (Almería, Spain) are described: Pygmar- rhopalites crepidinis Jordana & Baquero, n. sp. is characterized by trichobothria A, B and C forming a straight line towards posterior end; Ant. IV with five subsegments; all claws with inner tooth, all empodia with corner tooth, and empodial filaments surpassing tip of corresponding claw; anterior lobe of tenaculum with two apical chaetae; dens with 3, 2, 1,…, 1 anterior chaetae, and one exter- nal posterior spine. Pseudosinella najtae Jordana & Baquero, n. sp. is characterized by the absence of eyes; chaetotaxy R0R1R2011/10/0100+2, Abd. IV accessory chaeta s absent, all claws with four KEY WORDS teeth, paired teeth at different level (41 and 52% of claw length from basis), all empodia with ser- Arrhopalitidae, rate external edge, labrum with all chaetae ciliated; labium posterior without M1, with M2rEL1L2, Arthropoda, Pygmarrhopalites, r smooth microchaetae; remaining of labial chaetae (both posterior and anterior) ciliated. In addition Pseudosinella, one other species was found in the same cave: Troglopedetes absoloni (Bonet, 1931). The explorations Spain, of this cave have been carried out by the second author. The “Cueva del Saliente” is part of a study on biospeleology, seasonal changes, new cave fauna in Andalusia. In this cave a total of 4453 specimens (38 taxa) have been captured during species. an annual sampling period (October 2012 to August 2013).

ZOOSYSTEMA • 2017 • 39 (1) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 103 Jordana R. et al.

RÉSUMÉ Deux nouvelles espèces de collemboles (Hexapoda) de la grotte de Saliente (Almería, Espagne). Deux nouvelles espèces de collemboles cavernicoles appartenant aux genres Pygmarrhopalites Var- govitsh, 2009 et Pseudosinella Schäffer, 1897 et originaires de la « Cueva del Saliente » (Almería, Espagne) sont décrites : Pygmarrhopalites crepidinis Jordana & Baquero, n. sp. est caractérisée par des trichobothries A, B et C formant une ligne droite vers la partie postérieure ; Ant. IV avec cinq sous-segments; les griffes avec toutes une dent interne, tous les empodia avec une dent interne, et des filaments empodiaux dépassant la pointe de la griffe correspondante ; le lobe antérieur du tenaculum avec deux soies apicales ; la dens avec 3, 2, 1,… 1 soies antérieures, et une épine postérieure externe. Pseudosinella najtae Jordana & Baquero, n. sp. est caractérisée par l’absence d’yeux ; la chaetotaxie R0R1R2011/10/0100+2, Abd. IV sans soie s accessoire, toutes les griffes avec quatre dents, des MOTS CLÉS dents paires décalées (41 et 52 % de la longueur de la griffe en partant de la base), tous les empodia Arrhopalitidae, avec le bord externe dentelé, le labre avec toutes les soies ciliées ; labium postérieur dépourvu de M1, Arthropoda, Pygmarrhopalites, avec M2rEL1L2, microsoies r lisses ; autres soies labiales (tant postérieures qu’antérieures) ciliées. Pseudosinella, En outre, une autre espèce a été trouvée dans la même grotte : Troglopedetes absoloni (Bonet, 1931). Espagne, Les explorations de cette grotte ont été réalisées par le deuxième auteur. La « Cueva del Saliente » fait biospéléologie, changements saisonniers, partie d’une étude sur la faune cavernicole en Andalousie. Dans cette grotte un total de 4453 spéci- espèces nouvelles. mens (38 taxons) ont été capturés au cours d’un an d’échantillonnage (octobre 2012 à août 2013).

INTRODUCTION MATERIAL AND METHODS

Two cave projects have been developed along five years Cave description in order to protect invertebrate cave fauna in Andalusia The Saliente Cave is located in the mountains of the same (Belda et al. 2014). These projects were carried out in two name, on the hill of the Ermita, municipality of Oria, in periods: phase I (period 2009-2010) and phase II (2012- Almeria province (Spain). The entrance is located at 1170 m 2013), during which 32 caves were sampled in the eight a.s.l. UTM coordinates in 30S, X573269, Y4153731. The cav- Andalusian provinces. Six caves were sampled in Almeria ity is developed on alpujarrides carbonates in favor of a large province during the whole period and Saliente cave was area of fracture of NNW-SSE direction. In this area successive prospected during phase I. A wide variation has been fractures have slipped a large carbonated block that has broken observed in the abundance and ecological off the alignment which forms the sierra (González 2015). It structure between caves. The highest number of captures is a cavity where gravitational structures are dominant, with took place in the province of Cadiz during phase I, and in abundant chaos of blocks, diaclases, fractures and collapses. Malaga during phase II. There are superimposed galleries up to four levels. Its complex A total of 187 268 specimens of arthropods were processed structure, with spacious halls and labyrinthine galleries devel- and assigned to 104 taxonomic groups in a preliminary clas- oped among large detached blocks, makes the topographical sification. The material was sent to specialized taxonomists survey very complicated, which reaches 1190 m of accessible and it is under study. These projects are expected to yield at galleries, with maximum slope of – 56 m (Fig. 1). In the cavity least 33 new species, of wich 10 have been described already there is a large colony of bats, as well as an archaeological site. (Barranco 2010; Enghoff 2013; Molero et al. 2013; Ortuño & Barranco 2015; Ribera & De Más 2015). The material ob- Methods tained as part of these projects also has allowed the redescrip- The fauna was collected over two sampling periods. A first tion of rare forms and provided new data on known species preliminary sampling was conducted to determine the faunal (Ortuño & Barranco 2013; Gómez et al. 2013; Barranco potential capacity of the cavity (date setting of traps/collec- et al., 2014; Tinaut et al. 2015). tion date, 20.XI.-18.XII.2011). The following year we began The relevance of the taxonomic and faunistic results ob- a one-year study, placing traps at the stations established tained in the present study, has driven to the execution of during the preliminary sampling (28.X-01.XII.2012, 17.II-. some preservation measures for the protection of the fauna. III.2013; 17.III-21.IV.2013; 26.VI.-10.VIII.2013). In these These measures include, depending on each case, the instal- dates, 11 pitfall traps were installed throughout the cavity lation of fences around the entrance of the caves, to close (Fig. 1). Each trap consists of vials 120 ml and a diameter of the entrance cleaning of the caves and signaling of the ento- 5.2 cm mouth buried at the substrate level and filled up to mology importance of the cavern using informative boards half its capacity with a solution composed of ⅔ propylene (Belda et al. 2014). glycol and ⅓ beer solution, and primed with spicy sausage. In this paper, together with further information on the The traps remained active for about a month. The material fauna found in the “Cueva del Saliente”, two new species of extracted from each trap and that captured by hand was stored Collembola are described. in 70% ethyl alcohol.

104 ZOOSYSTEMA • 2017 • 39 (1) Fauna of Saliente cave

North wing South wing

North wing South wing The Hermitage T-02 Dolmen hall The Hermitage Entrance T-02 Dolmen hall T-03 T-Entrance10 T-11 T-09 T-10 T-03 T-05 T-11 T-04 Pintada hall T-09 SECTION T-01 T-06 T-04 T-05 Pintada hall SECTION Dolmen hall T-Enmedio01 hall T-06

T-08 Sinforosa hall Dolmen hall Enmedio hall Cuatro Escalones hall T-09 T-10 Pintada hall T-08 Sinforosa hall Cuatro Escalones hall T-10 T-09 Pintada hall T-07

–50 m T-07

–50 m PROFILES

The Hermitage PROFILES Sinforosa hall

The HermitageT-05 T-07 T-02 Sinforosa hall T-06 T-08 T-03 T-05 T-07 T-02 T-06 T-04 T-03 T-08

T-04

Cuatro Escalones hall

Dolmen hall T-01 Cuatro Escalones hall Enmedio hall Dolmen hall N T-01 T-10 Enmedio hall T-09 N –55.84 m T-10 Pintada hall

–50.85T-09 m –55.84 m T-11 Pintada hall

–50.85 m T-11

Fig. 1. — Topography of Cueva del Saliente and distribution of species of Collembola. Symbols: Star, Troglopedetes absoloni Bonet, 1931; triangle, Pseudosinella najtae Jordana & Baquero, n. sp.; square, Pygmarrhopalites crepidinis Jordana & Baquero, n. sp. T-1 to T-11 pitfall locations. Map base from the Asociación Espeleológica Velezana.

ZOOSYSTEMA • 2017 • 39 (1) 105 Jordana R. et al.

Relative humidity (RH) and temperature (T) were recorded SYSTEMATICS at hourly intervals during the sampling period by an Elec- tronic Dostmann LOG32 USB Data Logger located in the Class COLLEMBOLA Lubbock, 1870 Sinforosa Hall (traps 7 and 8, topography see Fig. 1). During Order Börner, 1901 this period the RH was 100% (ranging from 99.7-100%) Family Arrhopalitidae Stach, 1956 and the average T of 13.6°C (ranging from 13.5 to 13.7°C). Genus Pygmarrhopalites Vargovitsh, 2009 Some specimens were cleared in Nesbitt’s fluid and, after washing for one hour in 70% Ethyl alcohol, were mounted Pygmarrhopalites crepidinis Jordana & Baquero, n. sp. in Hoyer’s medium for compound microscope observation (Figs 2A-C; 3A-G, 4A-K; Table 1) in phase contrast and differential interface contrast (DIC). Some specimens were studied with a scanning electron Type material. — Holotype. ♀ in slide, date 21.IV.2013. Cueva microscope (SEM). Specimens from 70% ethyl alcohol del Saliente, Sierra de las Estancias, Cerro de la Ermita, Término were slowly rehydrated in decreasing series of its concentra- Municipal de Oria (Almería, Spain), pitfall 08, slide CIA-MZ- 20130421a, Barranco-ECA leg. tion (60%, 50%, 40%, 30%, 20%, 10%, distilled water) Paratypes. Barranco-ECA leg.: CIA-MZ-20130421d, 1 ♀ on slide, along 24 hours. Once in distilled water they were fixed in 4% 21.IV.2013, pitfall 08; MZ-20130421e, 1 ♀ on slide, 21.IV.2013, glutaraldehyde in cacodylate buffer for 24 h, and then trans- pitfall 08; MZ-20130421f, 1 ♀ on slide, 21.IV.2013, pitfall 08; MZ- ferred to sucrose 0.25 M for 24 h. The specimens were then 20130421g, 1 ♀ on slide, 21.IV.2013, pitfall 08; MZ-20130421h, 1 ♂ dehydrated up to 100% ethyl alcohol. Complete dessiccation (juvenile) on slide, 21.IV.2013, pitfall 07; MZ-20130421i, 1 speci- men on slide, 21.IV.2013, pitfall 09; MZ-20130421j, 1 specimen was achieved by the CO2 critical point technique. Samples juvenile on slide, 21.IV.2013, pitfall 02; CIA-MZ-20130421k and were then covered with a 16 nm thin layer of molecular gold l, 2 ♀ on SEM stub, 21.IV.2013, pitfall 08; MZ-20130810a, 1 ♀ using an Emitech K550 sputter coater. Observations were on slide, 10.VIII.2013, pitfall 05; MZ-20130810b, 1 ♀ on slide, done with a Zeiss Digital Scanning Microscope 940 A. The 10.VIII.2013, pitfall 05; MZ-20130317a, 1 ♀ on slide, 17.III.2013, chaetotaxy for Pygmarrhopalites Vargovitsh, 2009 follows pitfall 09; (deposited in MZ). CIA-MNHN-20130421b, 1 ♀ in slide, 21.IV.2013, pitfall 08, 3 in ethyl alcohol 70% (MNHN); Fjellberg (1998) for the labial palp, Fjellberg (1984) for CIA-MNCN-20130421c, 1 ♀ on slide, 21.IV.2013, pitfall 08, 3 in the outer maxillary palp, Christiansen & Bellinger (1996) ethyl alcohol 70% (MNCN); CIA-CECOUAL-20130317b, 2 ♀ on for antennal III sensory organ, Bretfeld (1999) for Abd. VI, the same slide, 17.IIV.2013, pitfall 08; CIA-CECOUAL-20130321, Christiansen (1966) and Christiansen & Bellinger (1980) 3 in ethyl alcohol 70% (CECOUAL). for empodium, and Vargovitsh (2009), for head, body and Complementary material. — Cueva del Saliente (Barranco- legs. The chaetotaxy for Pseudosinella follows Gisin & Da ECA/AEV leg): CIA-MZ-20111218a, 1 specimen juvenile in Gama 1969, Szeptycki (1979) and Soto-Adames (2010). slide, 18.XII.2011, pitfall 08; CIA-MZ-20111218b, 1 specimen The characters defined by Christiansen et al. (1990) for juvenile in slide, 18.XII.2011, pitfall 02; CIA-MZ-20111218c, Pseudosinella 1 ♀ on slide, 18.XII.2011, pitfall 07; CIA-MZ-20111218d, 1 ♀ and the characters used by Christiansen in on slide, 18.XII.2011, pitfall 09; CIA-MZ-20111218e, 1 specimen Jordana et al. (2016) in theirs Delta key have been used for juvenile on slide, 18.XII.2011, pitfall 08b; CIA-MZ-20111218f, identification and description. 1 ♀ on slide, 18.XII.2011, pitfall 04; CIA-MZ-20111218g, 1 ♀ on slide, 18.XII.2011, pitfall 07. Cueva del Saliente (Barranco-ECA Abreviations leg): CIA-MZ-20121201a, 1 specimen juvenile on slide, pitfall 07; Abd. abdomen or abdominal segment I-VI; CIA-MZ-20121201b, 1 ♀ on slide, 1.XII.2012, pitfall 06; CIA- Ant antennal or antenna/ae; MZ-20121201c, 1 ♀ on slide, 1.XII.2012, pitfall 05. CIA Cave Invertebrate Andalusia; Etymology. — The new species is named after Latin name of the DIC Differential Interference Contrast; cave: crepido, crepidinis. Mc Macrochaeta/ae; RH Relative humidity; Diagnosis. — Unpigmented; 1 + 1 eyes; trichobothria A, B and SEM Scanning Electron Microscope; T temperature; C forming a straight line towards hind part; Ant. IV subdivided Th Thorax or thoracic segments II-III. into five subsegments; all claws with inner tooth, all empodia with corner tooth, and empodial filaments overtopping tip of corresponding claw; anterior lobe of tenaculum with two Institutions apical chaetae; dens with 3, 2, 1…1 anterior chaetae, posterior CECOUAL Centro de Colecciones de la Universidad de Almería; MNHG Musée d’Histoire naturelle, Genève; side with one external spine; mucro gutter-like, both edges MNCN Museo Nacional de Ciencias Naturales, Madrid; serrated, tip rounded; some circumanal chaetae broadened MNHN Muséum national d’Histoire naturelle, Paris; with wings but in some specimens with a basal spine in a3 MZ Museum of Zoology, University of Navarra (Pamplona). without subbasal serration; anal appendices gutter-like, with apical and lateral end serration. RESULTS Description The only Collembola found in this cave are the two new spe- General cies and Troglopedetes absoloni Bonet, 1931, common in caves Body length 1.05 mm (n = 16) (holotype 1.22 mm), pigmen- of Andalusia, Spain (Soto-Adames et al. 2014). tation absent. Habitus as Figure 2A.

106 ZOOSYSTEMA • 2017 • 39 (1) Fauna of Saliente cave

A

B

C

DE

Fig. 2. — A-C, Pygmarrhopalites crepidinis Jordana & Baquero, n. sp.: A, habitus; B, end of dens and mucro; C, dorsal view of claw of leg 3; D, E, Pseudosinella najtae Jordana & Baquero, n. sp.: D, habitus, E, Labrum detail. Scale bars: A, 300 μm; B, 30 μm; C, 9 μm; D, 200 μm, E, 4 μm.

Female as in Figure 3E (Fjellberg 1998); maxillary outer lobe with Head (Fig. 3A, C). Length 0.34 mm (n = 18). Eyes 1 + 1. No three sublobal chaetae, and palp with basal spine (Fig. 3D) spine-like chaetae on the head.Clypeal area: 6 rows (from (Fjellberg 1984). a to f ); row a with axial chaeta, one axial chaeta between c, d and e rows. Inter-antennal area with 2 rows (α and β) Antennae (Fig. 4A-D). 2 times as long as head (n =16). without axial chaetae. Dorsal area: 4 rows (from A to D) Ant. length 0.70 mm (n = 16), ratio of Ant segments I : II : with 4 axial chaetae in rows A, B, C and D; B without lateral III : IV = 1 : 1.9 : 3.0 : 7.9 (average for n = 17). Ant. I with chaetae (Vargovitsh 2009). Labral chaetae number a: 4, m: 5, 7 chaetae, subapical posterior chaeta minute; Ant. II with p: 5; prelabral: 6 (Fig. 3C). Labial palp with papillae A-E 15 simple chaetae (Fig. 4A). Ant. III not swollen on sub-basal

ZOOSYSTEMA • 2017 • 39 (1) 107 Jordana R. et al.

D A C B A

β Th III f α B a p m m Th II e a 1 A d m c B b1 b a p C D p a c1 c2

a C pl

p m a

D F F3

p3 DL a 2 a1 1’ p3

F3 a1’ G DL a0 2 P3 DL E DL1 a0 1 a a a2’ 1’ 1 sa p1 a1 P1 a2’ sa’ a2 a3 sa’ sa a2 A1 A1 a 3 sa 2 VL1 av sa1 VL 1’ AV1 D 1 B AV1 sa1 av1’ VL2 1 sa3 av1’ 2 2 av5 3 AV2 av2 AV av4 3 2 2 3 VL6 p6 sa3 5 av3 AV3 6 VL VL5 A 7 av4 p VL4 6 C VL3 E

Fig. 3.— Pygmarrhopalites crepidinis Jordana & Baquero, n. sp.: A, head chaetotaxy, nomenclature for head chaetae rows: a-c and d, vertex; α and β, interanten- nal, a, b, c, d, e and f, clypeal area; pl, prelabral row; B, body chaetotaxy; C, labrum chaetotaxy (pl, prelabral row; p, posterior; m, medial; a, anterior); D, maxil- lary outer lobe; E, labial papillae A to E (posterior papillae drawn separately); F, Abd. VI chaetotaxy; G, schematic posterior view of Abd. VI. Scale bars: A, F, G, 50 μm; B, 100 μm; C, D, 20 μm; E, 10 μm. or medial part, with 17 chaetae and 2-rods sense organ; mi- D).Two species organites present and very patent on Ant. IV: crosensillum Aai, chaetae Api and Ap straight, chaetae Ape, subapical one with sphaeroidal big tip and apically special Ae and Ai curved (Fig. 4B). Ant. IV with 5 subsegments; curved guard sensillum (Fig. 4D) subsegmental formula: 1 + 3 + 1. AntIV bears the following whorls of chaetae: 5 on basal subsegment (BA, BM1-BM3, Legs (Fig. 4E-J). Foreleg: precoxae 1, 2 and coxa with 1, 0, 1 BB), 3 on medial subsegments each, and 5 on apical subseg- chaetae respectively; trochanter with 3 anterior and 1 posterior ment: AI-AIII, M1-M2 (after Vargovitsh, 2009) (Fig. 4C, chaetae; femur with 13 chaetae, a4 bending perpendicularly

108 ZOOSYSTEMA • 2017 • 39 (1) Fauna of Saliente cave

D I

C G

E

p i2

a i1

pi2 pi1

pe3 p3

ai1 a4

a1 p1

pe2

pe1 ae E, G, I 2 ae1 e1 C, D

K J FPpe FSa FH FPae FPe B FPpe FPpe Api Ap FPe FSa V Aai FSa FPe Ai Ape FPae Ae VV

IV Api A IV c IV

d III III III ab II II II

I I I

A, B

F, H, J

Fig. 4. — Pygmarrhopalites crepidinis Jordana & Baquero, n. sp. A, Ant. II; B, Ant. III; C, Ant. IV basal subsegment; D, Ant. IV distal four subsegments; E, subcoxae, coxa, trochanter and femur of leg 1, posterior view; F, tibiotarsus and claw of leg 1, posterior view; G, subcoxae, coxa, trochanter and femur of leg 2, anterior view; H, tibiotarsus and claw of leg 2, anterior view; I, subcoxae, coxa, trochanter and femur of leg 3, anterior view; J, tibiotarsus and claw of leg 3, anterior view; K, dens and mucro; a, b, c and d spines variations. Scale bars: 50 μm.

ZOOSYSTEMA • 2017 • 39 (1) 109 Jordana R. et al.

to the longitudinal axis of the segment (Fig. 4E) and pe3 Order ENTOMOBRYOMORPHA Börner, 1913, in a similar position; tibiotarsus (Fig. 4F) with 3 chaetae sensu Soto-Adames et al. (2008) FP (FPe, FPae, FPpe) and chaeta FSa present; whorl I Family Entomobryidae Tömösvary, 1882 with 9 chaetae; whorls II-V with 8 chaetae each. Pretarsus Subfamily Entomobryinae Schäffer, 1896 with 1 anterior and 1 posterior chaetae. Claw with inner Tribe Lepidocyrtini Yoshii & Suhardjono, 1989 tooth and 2 pairs of lateral teeth visible in ventral view Genus Pseudosinella Schäffer, 1897 (Fig. 4F), tunica absent. Empodium thin, with corner tooth in subbasal half, with long apical filament exceeding tip Pseudosinella najtae Jordana & Baquero, n. sp. of claw. Claw 4.5 times shorter than tibiotarsus. Mid leg: (Figs 2D, E; 5A-K) precoxae 1, 2 and coxa with 1, 1, 3 chaetae respectively; coxa with an additional spine; trochanter with 3 anterior Type material. — Holotype. ♀, slide CIA-MZ-20130421a, Cueva del Saliente, Sierra de las Estancias, Municipality of Oria, Almería, simple chaetae, 1 posterior chaeta and anterior trochanteral 21.IV.2013, pitfall 07, Barranco Leg. (as the remaining specimens organ; femur with 13 chaetae, 2 posterior ones smaller of type series). (Fig. 4G); tibiotarsus with 3 chaetae FP present, chaeta FSa Paratypes. CIA-MZ-20130421f to k, six specimens in ethyl alcohol, present; whorl I with 9 chaetae, whorls II-III with 8 chae- 21.IV.2013, pitfall 07; CIA-MZ-20130421l to u, 10 specimens in tae, whorl IV-V with 7 chaetae in different arrangement; ethyl alcohol, 21.IV.2013, pitfall 06; CIA-MZ-20130421l to m, two specimens mounted in SEM stub, 21.IV.2013, pitfall 07; CIA-MZ- claw broader than in foreleg, with inner tooth and 2 pairs 20130421b, ♀ in slide, 21.IV.2013, pitfall 06; CIA-MZ-20130421c, of lateral teeth (Fig. 4H), tunica absent; empodium broader juvenile in slide, 21.IV.2013, pitfall 06; CIA-MZ-20130810a, than in foreleg, with corner tooth in subbasal part; apical b and c, ♂, ♀ and juvenile in the same slide, 10.VIII.2013, pit- filament exceeding tip of claw; claw 5.5 times shorter than fall 06; CIA-MZ-20130810d, ♀ in slide, 10.VIII.2013, pitfall 06; tibiotarsus (Fig. 4H). Hind leg: precoxae with 1, 1 chaetae CIA-MZ-20130317b, juvenile in slide, 17.III.2013, pitfall 01; CIA-MZ-20130317c, juvenile in slide, 17.III.2013, pitfall 06 and coxa with 3 chaetae and 1 small spine; trochanter with (MZ). CIA-MNHN-20130421d, ♀ in slide, 21.IV.2013, pitfall 06 anterior trochanteral organ, 3 anterior and 1 posterior sim- (MNHN). CIA-CECOUAL-20130421e, ♀ in slide, 21.IV.2013, ple chaetae; femur with 10 anterior and 2 posterior chaetae pitfall 06 (CECOUAL). CIA-MNCN-20130317a, ♀ in slide, (Fig. 4I); tibiotarsus with 3 chaetae FP, chaeta FSa present; 17.VIII.2013, pitfall 04 (MNCN). whorls I-V as in mid tibiotarsus; claw broader than in fore Etymology. — The species name is dedicated to Judith Najt. and mid leg, with inner tooth and 2 pairs of lateral teeth (Fig. 4J); empodium broader than in the other legs, with Diagnosis. — Unpigmented; without eyes; chaetotaxy corner tooth in the middle, and apical filament exceeding R0R1R2011/10/0100 + 2, without s accessory chaetae on Abd. IV, all claws with four teeth, paired teeth at different level (50% of claw tip of claw; claw 7.5 times shorter than tibiotarsus (Fig. 4J). length), all empodia with serrate external edge, labrum with all chae- Length ratio of tibiotarsi I : II : III = 1 : 1.1 : 1.3. tae ciliated. Labium without M1, with M2rEL1L2, r is a smooth microchaetae, all labial chaetae both basal and internal ciliated. Great abdomen (Fig. 3A). Meso- and metathorax with normal dorsal chaetae, 1 neosminthuroid chaeta a on Th II Description and III. Trichobothria A, B and C forming a straight line General towards hind part. Single p chaeta of p-row of Abd. I is Maximum length 817 μm, mean length 727 μm (n = 8), located below the level of B trichobothrium, chaeta a1 (holotype 780 μm) (Fig. 2D). Without pigment. placed posterior to A trichobothrium, b1 placed between B and C trichobotria, chaeta c1 and c2 below C tricho- Head bothrium. Posterior lateral complex with 4 + 3 and furca Without eyes. Ratio antenna/cephalic diagonal 1.28, anten- base complex with 8 chaetae. Posterior dorsal complex with nal segments I/II/III/IV ratios 1/1.5/1.4/2.8 (average n = 7). about 23 chaetae arranged in 3 rows. Ventral complex with Sensorial chaetae s on antennal segment III short, slightly 3 chaetae. Tenaculum with 3 teeth, a basal peg-like projec- curved and rod like (Fig. 5D) similar sensillae are present in tion and 2 chaetae. external and lateral part of Ant. II and IV. Apical vesicle present on Ant. IV. Labrum chaetotaxy as in Figure 5A, all chaetae Furca. Manubrium with 5+5 posterior chaetae. Dens (Fig. 4K) ciliated (SEM microphotograph in Figure 2E). Formula of with 3, 2, 1,..,1 anterior chaetae, 3 inner, 6 dorsal and 7 ex- the labial base (Fig. 5B): MrEL1L2, all chaetae ciliated, r as ternal chaetae; two external spines with variable morphology smooth microchaetae. Antennae without scales. as Figure 4K (a, b, c) and 1 internal spine like chaeta (d). Body chaetotaxy Great abdomen. Abd. V with 2 chaetae and trichobothrium D Formula of the dorsal macrochaetae: R011/10/0100+2. R Mc in row a, and 3 chaetae in row p (Fig. 3B). Abd. VI (Fig. 3F, complex with R0, R1 and R2. S (M) as microchaeta, T and S G) without cuticular spines; some circumanal chaetae broad- present as Mc (Fig. 5C). Abd. II chaetotaxy: ‑aBq1q2 (Fig. 5J), ened with wings usually smooth, sometimes with weak distal without p (a2p) chaeta, a2i as ciliated microchaeta, a2 as serrations or basal spur (a1 and a3); chaeta A1 shorter than smooth microchaeta. B (m3) ciliated and long macrochaeta, chaeta a0 and DL2; subanal appendages (av5) gutter like, as in normal location, a3 near and external to as. q1 (a3e) and with apical serration. q2 (p4) as smooth microchaetae. Abd. III with a2i, a6-a6ii

110 ZOOSYSTEMA • 2017 • 39 (1) Fauna of Saliente cave

AB ED

C R0

R1

R2

S(M) F Tb

P(Pa5)0 T(S4)

a6i a m a6ii 2 as 3e a6 J a2i m5 K Ta 5 p psp a3 5 p p 6 m4 5p m3 p4 Tm2

a6i a6

a3 am6 a6ii Ta 5 G a2i a a7 2 m3 as m7a psp Tm5 m m7 4 m Tm2 6 a2ii p3 as p6 p7

B2 A1 B1 as A2 T1 B2p

F1 as D1 E1 H A3 B3 T2

C1 T3

E2 D2 T C2 4 F2 D2p E3

B5 T5 F2p A5 C3

B T6 A6 6 E4 I F3p C4

F4 T7 as D3

De3

Fig. 5. — A-E, G, I-K, Pseudosinella najtae Jordana & Baquero, n. sp.: labrum showing the ciliated chaetae (A); labium chaetotaxy (B); head chaetotaxy (the nomenclature follows Gisin & Da Gama 1969, and in brackets Soto-Adames (2010) (C); Ant. II–IV (D); claw (E); male genital complex (G); dens end and mucro (I); Abd. II–IV chaetotaxy (J); Abd. IV macrochaeta B5 (K); F, H, Pseudosinella infrequens Gisin & Da Gama, 1969 (drawings from holotype): claw and empodium (F); dens end and mucro (H). Scale bars: A, 4 μm; B, 15 μm; C, D, G, I, 20 μm; E, 10 μm.

ZOOSYSTEMA • 2017 • 39 (1) 111 Jordana R. et al.

Table 1. — Comparison among species that share the same characters in the claw complex (presence of corner tooth on claw, a tooth on internal lamella of ­empodium and a terminal filament on empodium surpassing the claw):A , dens, distal ventral or anterior chaeta (ve1): 0, normal; 1, strong spine; B, dens, ­external spine-like chaetae number; C, dens, E3 as spine-like: 0, normal; 1, spine-like; 2, strong articulated spine; D, dens, internal spine-like chaetae number; E, dens, L1 as spine-like: 0, normal; 1, spine-like; 2, strong articulated spine; F, dens, L2 as spine-like: 0, normal; 1, spine-like; G, dens, L3 as spine-like: 0, normal; 1, spine- like; H, annal appendix shape: 1, flat with apex and edges brush-like; 2, flat and pectinate;I , female anal valves, chaeta C1 on anal valve shape: 0, simple; 1, bi- furcate; J, female anal valves, chaeta C2 on anal valve: 0, normal; 1, swollen at the base; 2, winged; 3, dentate at the base; 4, branched; K, female anal valves, chaeta C3 on anal valve: 0, normal; 1, swollen at the base; 2, winged; 3, dentate at the base; L, female anal valves, chaeta C4 on anal valve: 0, normal; 1, swol- len at the base; 2, winged; 3, dentate at the base; 4, branched; M, female anal valves, chaeta C5 on anal valve: 0, normal; 1, swollen at the base; 2, winged; 3, dentate at the base; 4, branched; N, female anal valves, chaeta C6 on anal valve: 0, normal; 1, swollen at the base; 2, winged; O, female anal valves, chaeta C8 on anal valve: 0, normal; 1, swollen at the base; *, different; Dif, number of differences from the n. sp.

Species A B C D E F G H I J K L M N O Dif Pygmarrhopalites pygmaeus (Wankel, 1860) 0 2 2* 3* 1 1* 1* 1* 0 1* 1* 1* 1* 1 1 9 P. plethorasari Zeppelini, Taylor & Slay, 2009 – 2 2* 3* 1 1* 1* 1* 1* 1* 2 4 1* 1 1 8 P. lewisi Christiansen & Bellinger, 1998 0 2 2* 3* 1 ? ? 1* 0 1* 1* 1* 1* 1 0* 8 P. benitus (Folsom, 1896) 0 2 2* 3* 1 1* 1* 2* 0 1* 1* 1* 1* 1 1 9 P. ater Christiansen & Bellinger, 1998 0 1* 0* 1 1 0 0 2* 0 2 3* 3* 3* 2* 0* 8 P. maestrazgoensis Jordana, Fadrique & Baquero, 2012 0 1* 0* 0 0 0 0 1, 5?0 1* 0* 2 2 0(1) 0(1) 4 P. cantavetulae Jordana, Fadrique & Baquero, 2012 0 2 1 3* 2* 1* 1* 1, 5?0 3* 3* 3* 2 2* 0* 9 P. crepidinis Jordana & Baquero, n. sp. 0 2 1 0(1) 0(1) 0 0 5 0 2 2 2(4) 2 1 1 –

as cilliated microchaetae, there are three am6 microchaetae claw complex (presence of a tooth on claw, empodium with (two smooth and one ciliated) (Fig. 5J). Accessory chaeta s in a corner tooth on internal lamella and a terminal filament the anterior trichobothrial complex of abdominal tergite IV surpassing the claw) on all legs (Table 1). Among them only absent (Fig. 5J). P(C1) chaeta on abdominal tergite IV at level P. maestrazgoensis shares the external spines on the dens with of anterior tricobothrium (T2) and as smooth microchaeta. the new species. Table 1 only shows four differences, but the Only B5 and B6 presents as ciliated Mc as Figure 5K. Pseu- new species is also different by the chaetotaxy of Ant. organ III dopore at level of B5. Retinaculum with 4 + 4 teeth and one and Abd. VI (Jordana et al. 2012). ciliated chaeta. According to the dorsal macrochaetotaxy (R011/10/0100+2) of Pseudosinella najtae Jordana & Baquero n. sp. and the Legs absence of chaeta s in the anterior trichobothrial complex Claw (Fig. 5E) with 4 teeth, basal pair of wing teeth at 50% of the Abd. IV this species is similar only to Pseudosinella of the base of inner edge, and different in size, medial un- infrequens Gisin & Gama, 1969, described from a cave near paired tooth well developed, approximately 80% from base. Alhama de Murcia (Spain) from two specimens captured Empodium acuminate with serrate external edge. Tenent hair on bats’ guano. The holotype of P. infrequens (MHNG) has clavate, longer than claw. Scales only on coxae. been studied: it is different by some characters of the claw, i.e. three teeth (Fig. 5F) against four very patent teeth on Furca the internal edge of the claw of the new species (Gisin & Two internal and one external chaetae related to one distal Da Gama 1969: fig. 2), The empodium is serrated in both pseudopore of manubrial plate. Mucro with apical tooth species (not illustrated in the original description of P. infre- long as the apical; basal spine surpassing a little the tip of quens). Tenent hair a little longer than the claw in the new the anteapical tooth (Fig. 5I). Uncrenulated section of dens species and shorter in P. infrequens. Mucro of P. infrequens four times as long as mucro. Scales on manubrium and dens with apical tooth smaller than the subapical (Fig. 5H) draw- ing from the holotype. The labrum is very special in the new species with formula DISCUSSION 4/554 of prelabral/labral chaetae, which are all ciliated (Figs 2E; 5A), and with smooth labral chaetae on holotype. Labium In relation to Pygmarrhopalytes crepidinis Jordana & Baquero, without M1, with M2rEL1L2, r is a smooth microchaetae, n. sp. twenty species share the following characters: linear all labial chaetae both basal and internal ciliated, something bothriotrichal pattern in great abdomen head without spines unknown for P. infrequens (because the position the holotype and with 1+1 eye and Ant. IV with five segments. Among in the slide) in the original description only described as cili- them Pygmarrhopalites pygmaeus (Wankel, 1860) sensu Stach, ated the basal labial Mc. Both species are from caves of the 1918; P. benitus (Folsom, 1896) sensu Christiansen, 1966; Southern Spain: Alhama de Murcia vs Sierra de las Estancias, P. ater Christiansen & Bellinger, 1998; P. lewisi Christiansen & Municipality of Oria, Almería, which are 90 km apart. Bellinger, 1998; P. plethorasari Zeppelini, Taylor & Slay, 2009; Together with these two new species described above an P. maestrazgoensis Jordana, Fadrique & Baquero, 2012; P. can- additional accompanying fauna has been found. tavetulae Jordana, Fadrique & Baquero, 2012 and P. crepidinis The fauna in the cavity was analyzed based on 4468 speci- Jordana & Baquero, n. sp. share the same characters in the mens collected in the cave during the annual sampling period

112 ZOOSYSTEMA • 2017 • 39 (1) Fauna of Saliente cave

Table 2. — Fauna of Cueva del Saliente. Abbreviations: Coll, Collembola; Col, Coleoptera; Dip, Diptera; Hym, Hymenoptera; Pso, Psocoptera; Pse, Pseudoscor- pionida; Aca, Acari; Ara, Araneae; Iso, Isopoda; Chi, Chilopoda; Gas, Gasteropoda; Lep, Lepidoptera; AUT, Autumn; WIN, Winter; SPR, Spring; SUM, Summer.

Coll Col Dip Hym Pso Pse Aca Ara Iso Chi Gas Lep total T1 AUT 0 3 7 0 11 0 0 0 0 0 0 0 21 WIN 1 9 0 0 2 1 132 0 0 0 0 0 145 SPR 0 4 1 0 1 0 568 0 0 0 0 0 574 SUM 1 4 123 0 0 0 35 0 0 1 0 0 164 T2 AUT 2 2 4 0 0 0 0 0 1 1 0 0 10 WIN 0 1 4 0 0 0 0 0 1 0 0 0 6 SPR 1 1 4 0 0 0 0 0 0 3 0 0 9 SUM 0 1 6 0 0 0 0 0 0 0 0 0 7 T3 AUT 0 3 13 0 3 0 49 0 0 10 0 0 78 WIN 0 3 4 0 2 0 9 0 0 6 0 0 24 SPR 0 0 0 0 1 0 1 0 0 0 0 0 2 SUM 0 9 107 0 5 0 195 0 0 0 1 0 317 T4 AUT 0 18 43 0 0 0 2 0 0 8 0 1 72 WIN 2 0 8 0 11 0 97 0 0 0 0 0 118 SPR 0 5 11 0 3 0 46 0 0 0 0 0 65 SUM 0 3 71 0 4 1 50 0 0 3 0 0 132 T5 AUT 1 0 20 0 0 0 0 0 0 2 0 0 23 WIN 0 2 42 0 0 0 1 0 0 12 0 0 57 SPR 0 4 10 0 0 0 1 0 0 1 0 0 16 SUM 2 14 58 0 0 0 1 0 0 6 0 0 81 T6 AUT 1 55 94 0 0 0 0 0 10 10 0 0 170 WIN 1 0 1 0 0 1 3 0 0 1 0 0 7 SPR 18 0 0 0 0 0 0 0 0 2 0 0 20 SUM 4 1 28 0 1 0 2 1 0 0 0 0 37 T7 AUT 8 1 54 2 0 0 0 0 2 6 0 0 73 WIN 0 3 20 0 0 0 0 0 3 13 0 0 39 SPR 8 5 30 0 0 3 1 0 2 9 0 0 58 SUM 0 12 160 0 0 0 6 0 1 1 0 0 180 T8 AUT 0 15 139 1 0 0 0 0 26 5 0 0 186 WIN 68 1 26 0 0 0 0 0 0 6 0 0 101 SPR 64 4 54 0 0 0 1 0 0 1 0 0 124 SUM 1 63 113 0 0 1 10 0 0 9 0 0 197 T9 AUT 0 32 46 0 0 0 8 1 0 44 0 0 131 WIN 3 2 34 0 0 0 0 0 0 10 0 0 49 SPR 6 4 62 0 0 0 20 0 1 15 0 0 108 SUM 2 103 76 1 0 0 26 0 0 2 0 0 210 T10 AUT 0 8 225 0 0 0 21 0 0 0 0 0 254 WIN 0 5 7 0 0 0 6 0 0 28 0 0 46 SPR 0 7 11 0 0 0 4 0 0 7 0 0 29 SUM 0 0 86 0 0 0 26 0 0 3 0 0 115 T11 AUT 0 3 9 0 0 0 22 0 0 0 0 0 34 WIN 0 1 0 0 1 0 119 0 0 1 0 0 122 SPR 0 0 0 0 0 0 61 0 0 0 0 0 61 SUM 0 0 60 0 0 0 0 0 0 1 0 0 61 T12 AUT 0 4 7 0 8 0 115 0 0 1 0 0 135 Total AUT 12 144 661 3 22 0 217 1 39 87 0 1 1187 WIN 75 27 146 0 16 2 367 0 4 77 0 0 714 SPR 97 34 183 0 5 3 703 0 3 38 0 0 1066 SUM 10 210 888 1 10 2 351 1 1 26 1 0 1501 194 415 1878 4 53 7 1638 2 47 228 1 1 4468

(October 2012 to August 2013) (Table 2). All specimens have centage of specimens is Acari with 36.66%, the remaining been classified into 38 taxa with unequal taxonomic accuracy, taxonomic groups acount for less than 10% of specimens due the impossibility to identifying all the specimens to the (Table 2). Collembola represent 4.34%, with three species species level. All the material is being progressively determined (two of them new for science). to species level as taxonomic identifications by specialists in The three species of captured in this cave, have each group progress (Table 2). a different distribution pattern. Troglopedetes absoloni has The predominant group is the Diptera with 42.03% (con- been captured in the preliminary sampling and only in the sidering adults and larvae). The second largest group in per- pitfall 01, which was located very near the entrance, in the

ZOOSYSTEMA • 2017 • 39 (1) 113 Jordana R. et al.

twilight zone. It is probably an opportunistic or accidental Acknowledgements species in the cavity, since only three specimens, only in one This study was cofinanced by European FEADER funds and pitfall have been collected. Consejería de Medio Ambiente y Ordenación del Territorio The two species described in this work are unevenly de la Junta de Andalusia (Spain) via the Project “Propuesta de distributed in the cavity. Pseudosinella najtae Jordana & Servicio para la Conservación de los Invertebrados Amenazados Baquero, n. sp. has been captured from the entrance area to de Andalusia”. We thank to both caving clubs, Espeleo Club the pitfall number 07, but not beyond this point, in which a Almería (ECA) who made the sampling of the cave, the Aso- greater number of specimens have been captured. It would ciación Espeleológica Velezana (AEV) and Antonio González be risky to infer habits or ecological requirements of this Ramón who helped in samplings and provided the topography new species with only 32 specimens, but the area where of the cave. We thank the referees who have corrected this pa- the pitfall 06 was located is in the upper part of the cav- per and contributed to a substantial improvement. Also thank ity, where the environment is dry and dusty; environment Peter J. Schwendinger the Natural History Museum of Geneva and the room presents small isolated clusters of bat guano. for allowing observe the type of Pseudosinella infrequens. Pygmarrhopalites crepidinis Jordana & Baquero, n. sp. goes more deeply into the cavity, reaching pitfall number 09. Pit- fall 08 gave the largest number of captures. It is located in REFERENCES the Sinforosa room which is very active hydrologically and with abundant accumulation of guano. 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Submitted on 28 June 2016; accepted on 19 December 2016; published on 31st March 2017.

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